Sexual reproduction in the arum lily family, with emphasis on thermogenicity

Summary

Floral thermogenicity, which is found in several representatives of half a dozen angiosperm families, is most pronounced in the Araceae. It is based on the operation of an “alternative”, cyanide-resistant electron transport chain which, in contrast to the “classic” cytochrome oxidase system, produces little ATP; most of the energy originally locked up in the respiratory substrate usually starch — is therefore liberated in the form of heat. The biological function of this (biochemically wasteful) system is to release the heat to serve as a “volatilizer” for the floral odors (often containing aliphatic amines, indole and skatole) that attract the insect pollinators. This makes the survival value of thermogenesis (for the plant species) immediately clear. Thermogenicity is under tight biological control, as demonstrated by the fact that the same ceiling temperature is always reached, regardless of ambient temperature. In Eastern skunk cabbage (Symplocarpus foetidus), which flowers very early in spring, that ceiling is about 20° C, in tropical forms such as Xanthosoma robustum and Philodendron selloum, it lies in the 42°–44° C range. In several instances, e.g., in Arum and in Sauromatum, the voodoo lily, thermogenicity manifests itself as a flare-up of only a few hours' duration, a “respiratory explosion” that can lead to rates of metabolism that compare favorably with those of a hovering hummingbird. The metabolic peak is always reached at a particular time of day, which is different for the different arum lily species, and thus reduces competition for pollinators. The odors that accompany the heat are also very characteristic, appealing to different pollinator classes and further reducing such competition. In the voodoo lily and in Arum, the primary site for the production of both heat and odor is the naked appendix of the inflorescence, which acts as a specialized “osmophore” or odor carrier. The first explosion may be followed by another one several hours later, which manifests itself in the floral chamber of the inflorescence and is under strict photoperiodic control. In Sauromatum, the first metabolic explosion is triggered by a plant hormone, originally referred to as “calorigen,” which originates in the primordia of the staminate flowers and moves from there into the appendix where it exerts its action after a lag-time of about a day — an indication that synthesis of new enzymatic protein (through unblocking of certain genes?) may well be involved. In 1987, calorigen was shown to be identical with salicylic acid. This compound was already known to induce flowering in certain duck-weeds, Lemnaceae, which until recently were regarded as belonging to the same family as arum lilies. In certain water lilies (Nymphaeaceae), thermogenicity is combined with a pollination syndrome very similar to that of Arum and Sauromatum but involving temporary trap flowers.